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1.
F. Menu  D. Debouzie 《Oecologia》1993,93(3):367-373
Spreading of emergence over several years due to prolonged diapause in some larvae was shown in the chestnut weevil. Depending on the year the larvae buried themselves in the ground, 32–56% of live adults emerged after 2 or 3 years of underground life. Variability in the duration of diapause was assumed to correspond to tactics of adaptative coin-flipping plasticity. This plasticity must allow the chestnut weevil to respond to the unpredictability of its habitat as measured by the irregularity of chestnut production and summer drought. Indeed, fecundity and adult longevity did not lessen after 2 years of underground life. No drastic decrease in the population size of weevils occurs after bad years; for instance when the number of chestnuts on the study tree is less than 10 000, passers-by can collect all the fruit and about 95% of larvae developing in chestnuts are destroyed. Diapause nature (simple or prolonged) may be related to moisture and gas rates in the ground from October to December. These factors acting in autumn are not known to be involved in the physiological mechanisms that control the production of chestnuts.  相似文献   

2.
Within-generation variability in diapause duration can be viewed either as a mixed Evolutionary Stable Strategy (ESS), a genetic polymorphism of pure strategies, or as bet-hedging. Diapause variability expressed by a single genotype that maximizes mean geometric fitness at the cost of mean arithmetic fitness is a bet-hedging strategy. Bet-hedging differs from mixed ESS and stable genetic polymorphism of pure strategies because in these latter the expected pay-offs for all phenotypes are equal. In insects, individuals with a prolonged diapause (long cycle) lose at least one reproductive opportunity and suffer lower survival before reproduction than those with a short diapause (short cycle). If long-cycle individuals compensate this cost by better adult performance, the compensation leads to a trade-off which could result in mixed ESS or genetic polymorphism of pure strategies since the overall fitness of the two morphs may be similar. In this paper, we show that in the chestnut weevil Curculio elephas adult performance, measured as sex ratio, longevity, weight, and realized fecundity of females, are similar in individuals emerged after one and two years. Long-cycle morphs emerge slightly before short-cycle ones but this eventual advantage for fertility probably does not compensate higher larval mortality and missed reproductive opportunity in long-cycle phenotypes. Therefore, the cost associated with prolonged diapause cannot be completely compensated for by a better adult performance. From these results, and previous data, we conclude that variability in diapause duration cycle is better explained as bet-hedging than mixed ESS or genetic polymorphism of pure strategies.  相似文献   

3.
In many insect species, adult emergence spreads over several years because of the existence of prolonged diapause in certain individuals. From stochastic models, we show that diversified bet-hedging strategies (mixed strategies with emergence after 1 or 2 yr) are more fit than simple diapause strategy (emergence after 1 yr) or fixed prolonged diapause strategy (emergence after 2 yr) in isolated chestnut weevil populations. This conclusion applies to a large range of survival rates in prolonged diapause and is insensitive to initial conditions, magnitude of temporal autocorrelation, distribution of demographic parameters, and quoted values of population size limitation. However, the shape of the fitness distribution as a function of prolonged diapause frequency changes greatly in the absence of population size limitation. Whatever the survival rate during prolonged diapause, we find that there is no genotypic advantage to extending diapause for all chestnut weevil larvae to more than 1 yr. Our models predict selection of bet-hedging strategies over a large range of prolonged diapause frequencies. This result is consistent with the existence of several mixed strategies in a population. Emergences after 3 yr are not crucial for selection or for the dynamics of mixed strategies in the chestnut weevil.  相似文献   

4.
1. In some insects that overwinter as adults, mating occurs both before and after overwintering. Two hypotheses have been proposed to explain the adaptive significance of pre‐overwintering copulation of females. One is the bet‐hedging hypothesis, which explains pre‐overwintering copulation as a preparation for less chance of mating in the following spring. The other is the nuptial gift hypothesis, which states that secretions derived from males increase overwintering success of females. 2. In Eurema mandarina, both diapause autumn‐ and non‐diapause summer‐form male adults emerge with autumn‐form female adults in the last generation in a year. Most autumn‐form females mate with summer‐form males before winter, and re‐mate with autumn‐form males in the following spring. Because autumn‐form females have sufficient chances for mating after overwintering, the nuptial gift hypothesis has been regarded as the more probable hypothesis. 3. To test the nuptial gift hypothesis, the survival period was compared under short‐day conditions at 10 °C between mated and unmated females that had been reared on sucrose solution at 25 °C for 15–21 days. The mated females had significantly greater longevity than the unmated females, supporting the nuptial gift hypothesis. Body size also affected the survival period. 4. The results suggest that the nuptial gift is an important factor for the evolution of pre‐overwintering copulation in species in which females mate both before and after overwintering.  相似文献   

5.
The evolution of reproductive diapause is controversial in males as compared to females that must overwinter to leave offspring, because late‐autumn males can obtain offspring by pre‐overwintering copulation. The Japanese common grass yellow Eurema mandarina is suitable to examine the evolution of male reproductive diapause, because direct comparisons are possible between males that do and do not exhibit reproductive diapause. Approximately one‐half of males are insensitive to diapause‐inducing conditions, and emerge as non‐diapause summer‐form. Most autumn‐form females mate with summer‐form males in late autumn. Females that have overwintered re‐mate with autumn‐form males before the onset of oviposition. Because last‐male‐precedence is general in sperm competition in Lepidoptera, it is unclear why half of males emerge as summer‐form in late autumn. A potential adaptive benefit for emerging as summer‐form is increased sperm overwintering success, if autumn‐form females have a higher overwintering success than autumn‐form males. In the present study, overwintering success was estimated for both sexes of autumn‐form adults by rearing under seminatural conditions and a mark–release–recapture technique. Both approaches estimated an overwintering success of approximately 5% for both sexes. The absence of difference in overwintering success between the sexes suggests that pre‐overwintering copulation does not increase sperm overwintering success. However, a considerably low overwintering success may explain, at least partly, the presence of summer‐form males in late autumn. The degree of overwintering success might be more important than the sexual differences of overwintering success in the evolution of male reproductive diapause.  相似文献   

6.
Abstract Diapausing insect species have evolved a great diversity of life cycles, although overwintering occurs at a single development stage within most species. Understanding why diapause has evolved towards a given life stage requires investigation of both the ecological and physiological attributes. Notably, it is suggested that adult overwintering is more energy‐demanding than larval overwintering but it brings fitness gains by allowing adults to be synchronized with their seasonal requisites through an early spring emergence. This hypothesis is tested in field conditions in the nut weevil Curculio nucum, whose life cycle comprises an obligate 2‐year, nonfeeding underground phase, including a larval, followed by an adult, overwintering. In this species, adult wintering leads to an early spring emergence; at first glance, however, this does not enhance synchronization between weevils and their host because adults emerge more than 1 month before starting to breed. It is suggested that adult overwintering ultimately evolved in response to the phenology of the host, by allowing females to oviposit in nuts before their full sclerotinization. Adult overwintering appears to be costly because adults postpone reproduction for 1 year, incur a significant weight loss and require feeding before egg laying. Surprisingly, lipids are unaffected during diapause, lipogenesis even being likely in the summer metamorphosis. These results suggest that the lipids involved in egg production may entirely come from the larval stages, whereas the other nutrients are acquired through adult feeding but this remains to be tested.  相似文献   

7.
Extended life cycle in the chestnut weevil: prolonged or repeated diapause?   总被引:2,自引:0,他引:2  
Many insect species extend the life cycle of a part of their population over several years. The adaptive value of these long cycles is now well documented, but the physiological processes underlying them have been little studied. Long life cycles are usually viewed as resulting from prolonged diapause proceeding from a simple extension of the usual winter diapause. However, this hypothesis has not been greatly tested, and information is lacking for species with a larval diapause. The energetic cost of a prolonged larval diapause also needs to be measured, because the few published estimates of lipid consumption concern an imaginal diapause. It is not therefore clear whether the negligible lipid consumption observed during adult prolonged diapause can be extrapolated to larval diapause. From microrespirometry and lipid measures in the chestnut weevil, Curculio elephas Gyllenhal (Coleoptera: Curculionidae), we show that: (1) in contrast to the usual hypothesis, the long cycle does not result from an extension of larval winter diapause, but is due to a prolonged diapause occurring secondarily to a developmental phase, and (2) energy consumption during the prolonged diapause is not negligible, but is provided for by a higher initial lipid content in the long cycle individuals. The adaptive value of the observed cycle is discussed.  相似文献   

8.
对浙江省双季稻区稻水象甲(LisorhoptrusoryzophilusKuschel)夏季种群的研究表明,光照和温度可排除在一代成虫夏季滞育的诱导因子之外,而不同的食料条件是引发滞育的主导因素.在自然状态下,即使是强迫滞留,早稻田内的一代成虫也不转入正常发育,但取食稻秧能使新羽一代成虫和稻田内外处于生殖滞育状态的一代成虫的卵巢正常发育,并能解除已进入夏蛰的一代成虫的生殖滞育;同时,不论是新羽成虫还是夏蛰成虫,取食稻秧后卵巢和飞行肌同时发育;若这种食料条件能持续下去,则飞行肌消解而卵巢发育产卵.  相似文献   

9.
Cephalcia arvensis Panzer (Hym., Pamphiliidae) spends most of the life cycle as a prepupa inside an earth-walled chamber in the soil of spruce (Picea abies Karst.) forests. There are two types of prepupae in diapause in the soil, the eonymph and the pronymph. Prolonged diapause as an eonymph occurs in prepupae just after soil penetration at temperatures below 12°C. Eonymphs change into pronymphs during the following summer and the life cycle lasts for two years. Mature larvae that enter the soil at temperatures above 12°C change immediately into pronymphs and these individuals will emerge during the next spring. Other factors such as photoperiod do not appear to be correlated with prolonged diapause. The depth reached by the mature larvae in the soil is proportional to their size and inversely proportional to the soil temperature. Consequently, voltinism depends on both the conditions affecting development of the larvae in the crowns and the temperature at different soil depths.  相似文献   

10.
Frequency distributions of insect immatures per host are often fitted to contagious distributions, such as the negative binomial, to deduce oviposition pattern. However, different mechanisms can be involved for each theoretical distribution and additional biological information is needed to correctly interpret the fits. We chose the chestnut weevil Curculio elephas, a pest of the European chestnut Castanea sativa, as a model to illustrate the difficulties of inferring oviposition pattern from fits to theoretical distributions and from the variance/mean ratio. From field studies over 13–16 years, we show that 20 out of the 31 yearly distributions available fit a negative binomial and 25 a zero-inflated Poisson (ZIP). No distribution fits a Poisson distribution. The ZIP distribution assumes heterogeneity within the fruit population. There are two categories of host: the first comprises chestnuts unsuitable for weevil oviposition or in excess relative to the number of weevil females, and the second comprises suitable fruits in which oviposition behavior is random. Our results confirm this host heterogeneity. According to the ZIP distribution, the first category of hosts includes on average 74% of the chestnuts. A negative binomial distribution may be generated by either true or false contagion. We show that neither interference between weevil females, nor spatial variation in the infestation rate exist. Consequently, the observed distributions of immatures are not the result of false contagion. Nevertheless, we cannot totally exlude true contagion of immatures. In this paper we discuss the difficulty of testing true contagion in natural conditions. These results show that we cannot systematically conclude in favour of contagion when fitting a distribution such as the negative binomial or when a variance/mean ratio is higher than unity. Received: 22 September 1997 / Accepted: 15 December 1997  相似文献   

11.
Zamia furfuracea, a cycad of Mexican origin, in cultivation is pollinated by the snout weevil, Rhopalotria mollis, also of Mexican origin. This weevil apparently is host specific, swarming upon male cones of the cycad, where mating, feeding, and oviposition occur. Sporophylls of male cones are rich in starch; those of female cones are poor in starch, and weevils feed upon male cones and are visitors to but not feeders upon or within female cones. Pollen transport to female cones occurs during such visitation. All stages of metamorphosis of R. mollis occur within male cones; larvae feed exclusively on parenchyma of microsporophylls, pupate within stalks of microsporophylls, and emerge as adults from the outer ends of microsporophylls. They do not feed on pollen and do not damage microsporangia or pollen. Toward the end of the breeding season of the weevil (and the cycad), some larvae enter diapause in thick-walled pupal cases within microsporangial stalks of pollen-spent cones. These remain in diapause until the next reproductive season of the cycad.  相似文献   

12.
This study examines whether in nature endangered quino checkerspot (Euphydryas editha quino) larvae will return to diapause and if so where they choose to hide. Multiple years of diapause probably help larvae survive drought years and sites chosen have high survival value to the species. Ninety square meters of habitat were created by removing non native plants and replacing them with natives found at checkerspot occupied sites. During the 2005–2006 winter 1,000 post-diapause larvae were released. From these larvae 31 adults (20 males and 11 females) developed over a 2.5 month period (March 20–June 6) from 41 pupae. One chrysalis was parasitized by a parasitic wasp Pteromalus puparum (L.) in the family Pteromalidae, one was partially eaten by an animal, while the remaining eight pupae died of unknown causes. Thirty quadrats (1 square meter each) were cleared of vegetation, leaf and branch litter, rocks, and checkerspot larvae from July 5 to August 1, 2006. Forty-nine larvae were found that returned to diapause. Most larvae (31) chose to make shelters on California buckwheat, which is not a checkerspot food plant, two to five cm above the ground. One shelter had 22, another had seven, and two others had single larvae. Five of 10 larvae found in leaf litter below California buckwheat were crawling and not associated with shelters suggesting they had been dislodged from shelters. California buckwheat may be important in habitat restoration for the checkerspot, particularly at sites below 900 meters elevation where summer conditions are hot and dry. No additional larvae were found the following spring, when they should have exited diapause. Therefore 910 (91%) larvae were lost to some undocumented form of mortality.  相似文献   

13.
The introduced African dung beetle, Onitis alexis Klug, has become established in the warmer regions of Australia. The south-eastern limit of its current distribution is Moruya, NSW, and the Araluen Valley 50 km inland. At Araluen newly emerged beetles are present in dung in late spring, summer and autumn. Egg-laying starts 1-2 weeks after emergence and continues throughout the summer and autumn, as indicated by the presence of parous females in the population and of broods under experimental pads. Eggs laid in December/January produce adults in late summer and autumn, those laid from February to April produce adults in the following spring and summer. In the laboratory, mortality of larvae is high in cold (0–16°C), wet conditions and their development is delayed in warm (25°C and 27°C), dry conditions. This delay was confirmed in the field during the summer drought of 1982-83 when predicted times of emergence (based on day-degree summation in the soil) always preceded the observed emergence time of the local population, as well as preceding the emergence of beetles developing from eggs laid at known times. Follicle resorption in adult females was related directly to increasing age and to rainfall. Dung collected from hayed-off pasture did not affect fecundity, but caused larval mortality. Adults survived the winters at Araluen in some years, and immatures survived best during dry winters, being facilitated in this by a cold-induced larval diapause. Onitis alexis larvae can survive wet or dry summers, and cold dry winters (down to about 0°C) but not wet winters. This seems to be the major factor limiting the southern distribution of the species.  相似文献   

14.
Abstract. Galerucella nymphaeae L. , a chrysomelid that feeds on Lythrum spp., water lily and water chestnut, is closely related to two European species that were recently introduced into North America for biological control of L.salicaria , purple loosestrife. To develop a paradigm for continuously rearing these and other univoltine chrysomelids, we conducted field and laboratory studies on G. nymphaeae's development , reproduction and diapause.
A high incidence of reproduction without diapause occurred when fourth instars, pupae, and adults (held as pairs) experienced very long daylengths (LD18:6 h), i.e. longer than those G.nymphaeae encounters in nature. Under similar photoperiodic conditions, adults maintained in groups showed a significantly higher rate of reproductive diapause than those held as pairs. Females laid three to seven egg masses/week, and the size of egg masses varied between nine and nineteen eggs. During their reproductive lifetimes, individual females showed a highly significant propensity to lay a consistent number of eggs/egg mass.
After diapausing under short daylengths and low temperature (LD 10:14h, 5C), adults transferred to long days (LD 18:6h at 21C) had high rates of diapause termination and postdiapause oviposition. In contrast, those transferred to short daylengths (LD 10:14h at 21C) had low rates of reproduction.
Laboratory-derived heat-degree models accurately predicted egg and pupal, but not larval, development in the field. In nature, most females in the summer generation entered reproductive diapause without ovipositing; a small proportion of females that emerged relatively early (by mid June) oviposited before entering diapause. The overwintering population consists of adults from the first-generation and a small number from the second generation.  相似文献   

15.
Abstract The white grub Dasylepida ishigakiensis has a 2‐year life cycle and spends approximately 9 months as a nonfeeding larva, pupa and adult on a subtropical island. Evidence is presented indicating that this beetle has two diapauses that appear to synchronize this long life cycle with the seasons. Larvae exposed to 20, 22.5, 25 and 27.5 °C late in the third (last) stadium pupate rapidly except for some individuals kept at the highest temperature. The latter pupate upon transfer to 22.5 °C, indicating that larval diapause is maintained at high temperature but terminates upon transfer to a lower temperature. Pupal development is directly temperature‐dependent in the range 20–30 °C. Adults develop reproductive organs (i.e. the ovary in females and the seminal vesicles and accessory glands in males) rapidly at 15 and 20 °C, whereas those kept at 25 °C take a long time to do so. Ovarian development is completely suppressed at 30 °C but initiated upon transfer to 20 °C. In the laboratory, males with well‐developed reproductive organs mate even with sexually immature females , whereas females with undeveloped ovaries show no sexual behaviour. Although the two diapauses of this species are thermally regulated (i.e. a characteristic commonly expressed by insects in summer diapause), adults of this beetle emerge from pupae late in the autumn and remain in the soil for 2 months. Adult diapause effectively serves to synchronize the time of sexual maturation with the coldest month of the year.  相似文献   

16.
The chestnut weevil Curculio sikkimensis undergoes a prolonged larval diapause that is completed by repeated exposure to chilling and warming. We examined the possible reversibility of diapause intensity in response to temperature changes. All larvae were subjected to an initial chilling followed by incubation at 20°C to force pupation of the 1-year-type larvae that require only one winter for diapause completion. We then exposed the larvae remaining in prolonged diapause to a second chilling at 5°C for different lengths of time, preceded or not preceded by incubation at 20°C (moderately high) and/or 25°C (high) and followed by a final post-chilling reincubation at 20°C. Many of the prolonged-diapausing larvae subjected only to a brief second chilling were re-activated upon reincubation. However, short exposure to 25°C before this second chilling dramatically decreased the percentage of larvae completing diapause. When larvae were exposed to 25°C for a short period, then incubated at 20°C and subjected to the brief second chilling, many were re-activated during reincubation. The chilling time required for most of the larvae to complete diapause decreased after pre-chilling incubation at 20°C and increased after incubation at 25°C. These results demonstrate that diapause intensity in C. sikkimensis changes reversibly in response to changes in ambient temperature.  相似文献   

17.
《Acta Oecologica》2002,23(1):31-39
The chestnut weevil, Curculio elephas (Gyll.), is a non-outbreaking species whose populations and food resources, the European chestnut, Castanea sativa, can be precisely defined. Thirteen and 17 generations of this insect were studied in two isolated sites. Field observations and experiments allowed us to estimate the absolute abundance, availability and use of chestnuts for weevil oviposition, and the number of weevil females emerging per site. Unavailable chestnuts were defined as the fruits either infested first by the chestnut moth (Cydia splendana) larvae (because of competition between the two species) or those avoided by chestnut weevil females when selecting their egg-laying sites, independently of chestnut moth presence. From a third to a half of the chestnuts were not available on the average for weevil infestation. Only one-fourth, on the average, of those available for oviposition were actually used by chestnut weevil females. Regardless of year and site, the number of available chestnuts per weevil female was higher than that of weevil-infested fruits per female, considering global food resources independently of their temporal variation in quality. However, realized fecundity of weevil females was positively correlated with the mean number of available chestnuts per female. We concluded that food resources can be limiting without being fully exploited by females because of temporal variation in chestnut quality.  相似文献   

18.
Adults of the chestnut weevil Curculio sikkimensis emerged over a 3-year period under laboratory and quasi-field conditions due to a prolonged diapause that occurred at the mature larval stage. Variable proportions of the larvae remained in diapause after a single cold (5 degrees C) treatment of 120 days. Extension of the chilling period to as long as 540 days did not increase the percentage of diapause termination, and excessively long chilling actually reduced the percentage. Chilling was not indispensable to the termination of larval diapause. Diapause intensity was very high and variable, and more than 1000 days at 20 degrees C was necessary to reactivate all diapause larvae. When the diapause larvae were exposed to cycles of low (5 degrees C for 120 days) and high (20 degrees C for 240 days) temperatures, the percentage of diapause termination reached 100% after two or three such cycles. Thus, the prolonged diapause of C. sikkimensis has characteristics similar to the common short winter diapause in other insects, but has unique characteristics that ensure polymodal reactivation over several years.  相似文献   

19.
Three methods of short-term storage of the blowfly Calliphora vicina strains are considered based on the experimental study of 21 strains originating from different parts of the species range. The colony can be preserved as diapausing adults at 6° and darkness for 2–3 months or more, depending on the geographical origin of the population. During the first five days of adult life the flies should be kept at 12° and short day on a sugar diet, after which they should be transferred into a refrigerator. During artificial hibernation the flies also require periodic sugar feeding every 20 days (3–4 h at 20°C) to maintain their vital functions. The combination of temperatures of 20–23°C and a protein diet terminates reproductive diapause, and oviposition starts in 10–17 days. The fly strain may be preserved as reproductive females at 6°C and darkness with sugar feeding. Flies also require periodic sugar feeding at 20°C (3–4 hours). In this case the flies start laying eggs 2–3 days after being transferred to 20–23°C. The preservation of diapausing larvae is a more reliable method of prolonged strain storage. In this case the flies of maternal generation are maintained at 20–23°C on sugar and protein diet. The egg rafts laid during 5–6 hours are then transferred into 12°C and short day until hatchment. The hatched larvae should be immediately placed into a refrigerator (2–3 or 5–6°C), where they feed during 1–1.5 months and enter diapause. For strain restoration, the diapausing larvae should be transferred into 20–25°C, where they pupariate in 3–5 days and the flies emerge in nearly 10 days.  相似文献   

20.
Plants in the Mediterranean climate region of California typically experience summer drought conditions, but correlations between zones of frequent coastal fog inundation and certain species’ distributions suggest that water inputs from fog may influence species composition in coastal habitats. We sampled the stable H and O isotope ratios of water in non-photosynthetic plant tissue from a variety of perennial grass species and soil in four sites in northern California in order to determine the proportion of water deriving from winter rains and fog during the summer. The relationship between H and O stable isotopes from our sample sites fell to the right of the local meteoric water line (LMWL) during the summer drought, providing evidence that evaporation of water from the soil had taken place prior to the uptake of water by vegetation. We developed a novel method to infer the isotope values of water before it was subjected to evaporation in which we used experimental data to calculate the slope of the δH versus δO line versus the LMWL. After accounting for evaporation, we then used a two-source mixing model to evaluate plant usage of fog water. The model indicated that 28–66% of the water taken up by plants via roots during the summer drought came from fog rather than residual soil water from winter rain. Fog use decreased as distance from the coast increased, and there were significant differences among species in the use of fog. Rather than consistent differences in fog use by species whose distributions are limited to the coast versus those with broader distributions, species responded individualistically to summer fog. We conclude that fogwater inputs can mitigate the summer drought in coastal California for many species, likely giving an advantage to species that can use it over species that cannot.  相似文献   

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